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Rivista Italiana di Paleontologia e Stratigrafia (Research in Paleontology and Stratigraphy) vol. 125(3): 587-608. November 2019 RECENT BRACHIOPODS FROM THE TONGA ISLANDS, SW PACIFIC: TAXONOMY AND BIOGEOGRAPHY MARIA ALEKSANDRA BITNER Institute of Paleobiology, Polish Academy of Sciences, ul. Twarda 51/55, 00-818 Warszawa, Poland. E-mail: [email protected] To cite this article: Bitner M.A. (2019) - Recent brachiopods from the Tonga Islands, SW Pacific: taxonomy and biogeography. Riv. It. Paleont. Strat., 125(3): 587-608. Keywords: Brachiopoda; systematics; biodiversity; Pacific archipelagos; BORDAU 2. Abstract. Twenty species of Recent brachiopods belonging to the genera Neoancistrocrania, Basiliola, Basiliolella, Dyscolia, Abyssothyris, Xenobrochus, Terebratulina, Fallax, Septicollarina, Frenulina, Amphithyris, Annuloplatidia, Leptothyrella, Dallina, Campages, Thecidellina and Minutella have been identified in the material collected during the French cruise BORDAU 2 to the Tonga Islands, South-West Pacific. Apart from Frenulina sanguinolenta all species represent the first records for the Tonga Islands. The investigated brachiopod fauna shows the greatest affinity to that from Fiji and New Caledonia, having 16 and 12 species in common, respectively. Although less affinity is observed with the New Zealand fauna, there are two species, Terebratulina australis and Amphithyris buckmani reported so far only from New Zealand, Fiji and Tonga. The biodiversity of brachiopods in Tonga is similar to that in Fiji but half as great as that in New Caledonia and New Zealand regions and much higher than in French Polynesia. INTRODUCTION origin, the eastern non-volcanic coral limestone. So far only two species, Novocrania turbinata (Poli, 1795) Since 1976 the South-West Pacific region has and Frenulina sanguinolenta (Gmelin, 1791) have been been intensively surveyed within the program Trop- reported from Tonga (Saito & Endo 2001; Logan ical Deep-Sea Benthos (formerly MUSORSTOM) 2007; Robinson 2017). Although both species are established by the Muséum national d’Histoire na- well known and widely distributed in the western turelle (Paris, France) and the Institut de Recher- Pacific, in the studied material they are either absent che pour le Développement (Nouméa, New Cale- (N. turbinata), or very rare (F. sanguinolenta). How- donia). The present report focuses on the Tonga ever, Robinson (2017) recognized N. turbinata in the island group, explored during the cruise BORDAU 2 University of Tokyo collection from Tonga. Among in 2000. The name of the cruise refers to the BOR- the 20 species identified in the studied material, 19 Der of the Indo-AUstralian plate (Bouchet et al. are reported for the first time from the Tonga Is- 2008). Tonga, comprising more than 170 islands, lands. is an archipelago in the South Pacific Ocean close to Fiji, about two-thirds of the way from Hawaii to New Zealand (Fig. 1). It lies along the boundary of the Pacific and Indian-Australian tectonic plates MATERIAL AND METHODS and consists of two geologically different, parallel The material presented here was collected during the cruise chains of islands; the western islands are of volcanic Bordau 2 (https://expeditions.mnhn.fr/campaign/bordau2) to the Tonga Islands, SW Pacific (Fig. 1). The expedition was organized by the Muséum national d’Histoire naturelle, Paris and by the Institut Received: March 12, 2019; accepted: May 17, 2019 de la Recherche pour le Développement, Nouméa, New Caledonia 588 Bitner M.A. Class CRANIATA Williams, Carlson, Brunton, Holmer & Popov, 1996 Order Craniida Waagen, 1885 Superfamily Cranioidea Menke, 1828 Family Craniidae Menke, 1828 Genus Neoancistrocrania Laurin, 1992 Type species - Neoancistrocrania norfolki Laurin, 1992, by original designation of Laurin (1992: 344) Neoancistrocrania norfolki Laurin, 1992 Fig. 2A-C 1992 Neoancistrocrania norfolki Laurin, pp. 344-346, pl. 1, figs. 1-6, pl. 2, figs. 1-6. 1997 Neoancistrocrania norfolki - Laurin, pp. 417-418, fig. 41A-E. 2009 Neoancistrocrania norfolki - Bitner, p. 6, fig. 2A-F. 2010 Neoancistrocrania norfolki - Zezina, p. 1179. 2014a Neoancistrocrania norfolki - Robinson, fig. 3A-H. 2014b Neoancistrocrania norfolki - Robinson, p. 542, fig. 2A-E. 2015 Neoancistrocrania norfolki - Bitner, p. 35, fig. 2A-B. Material examined: Tonga, BORDAU 2 cruise, stn DW 1535, one dorsal valve (MNHN IB-2013-678); stn DW 1602, one dorsal valve (MNHN IB-2013-679); stn DW 1605, one ventral valve, broken (MNHN IB-2013-680). Depth range: 263-441 m. Measurements: Max. length 14.8 mm, width 20.8 mm. Remarks. Neoancistrocrania norfolki is very rare in the studied material, found in only 3 stations. Morphologically this species is readily distinguished Fig. 1 - Map of Tonga Islands, showing location of the brachio- from Novocrania Lee & Brunton, 2001 by its massive, pod-bearing stations. mineralized ventral valve and two erect divergent processes on the dorsal valve interior (Laurin 1992, on R.V. “Alis” and carried out from 31 May 2000 to 22 June 2000. 1997; Bitner 2009, 2015) and by unique features of Brachiopods were found in 69 out of 138 stations. The exact loca- tion, depth and species identified at each stations are given in the the soft tissues (Robinson 2014a, 2014b). However, Appendix. Although the collection is rich overall (1212 specimens), molecular analysis shows that “Neoancistrocrania is nine species are rare (3 or fewer specimens). no more distantly related to the Northern clade of Samples were collected using a Warén dredge (DW) or a Novocrania than are other Novocrania clades” (Cohen trawl (CP, CH). To remove soft tissues, specimens were treated with hypochlorite bleach, followed by a water wash. For scanning elec- et al. 2014: 145) suggesting generic synonymy of tron microscope (SEM) examination, the selected specimens were Neoancistrocrania under Novocrania. Such conflict be- mounted on stubs, coated with platinum, and investigated using a tween the morpho-classification and molecular sys- Philips XL-20 microscope at the SEM laboratory of the Institute of tematics is observed in other brachiopod groups as Paleobiology, Warszawa. All the material is deposited in the collection of the Museum national d’Histoire naturelle, Paris, France under the well (Cohen & Bitner 2013; Bitner & Cohen 2015). catalogue number MNHN IB-2009-516 to IB-2009-562, IB-2009- Previously N. norfolki had been found only in 1198, IB-2009-1240, IB-2013-678 to IB-2013-790. the West Pacific (Laurin 1997; Bitner 2009, 2015; Cohen et al. 2008, 2014). The present finding ex- tends its geographical range eastward. SYSTEMATIC PART Subphylum RHYNCHONELLIFORMEA Phylum BRACHIOPODA Duméril, 1805 Williams, Carlson, Brunton, Holmer & Popov, 1996 Subphylum CRANIIFORMEA Popov, Bassett, Class RHYNCHONELLATA Williams, Carlson, Holmer & Laurie, 1993 Brunton, Holmer & Popov, 1996 Recent brachiopods from the Tonga Islands, SW Pacific 589 Fig. 2 - A-C - Neoancistrocrania norfolki Laurin, 1992, Tonga, BORDAU 2, stn DW 1602, 263-320 m, outer, inner and tilted views of dorsal valve, MNHN IB-2013-679. D-H - Basiliola lucida (Gould, 1862); (D-E) inner and oblique views of posterior part of dorsal valve, MNHN IB-2013-700, SEM, stn DW 1635, 320-323 m; (F-H) dorsal, lateral and anterior views of articulated specimen, MNHN IB-2013-699, stn DW 1587, 309-400 m. I-M - Basiliola beecheri (Dall, 1895), BORDAU 2; (I-K) dorsal, lateral and anterior views of articulated specimen, MHNH IB-2013-694, stn DW 1619, 591-593 m; (L-M) inner and oblique views of posterior part of dorsal valve, MNHN IB-2013- 683, SEM, stn CP 1545, 444-447 m. N-S - Basiliolella colurnus (Hedley, 1905), BORDAU 2; (N-Q) dorsal, lateral views and dorsal, anterior views of two articulated specimens, MNHN IB-2013-706, stn CP 1545, 444-447 m; (R-S) inner and oblique views of posterior part of dorsal valve, MNHN IB-2013-703, SEM, stn DW 1536, 320-323 m. 590 Bitner M.A. Order Rhynchonellida Kuhn, 1949 1895 Hemithyris beecheri Dall, p. 717, pl. 31, figs. 1-4. 2008 Basiliola beecheri - Bitner, pp. 427-428, fig. 5H-L cum( syn.). Superfamily Pugnacoidea Rzhonsnitskaya, 1956 2009 Basiliola beecheri - Bitner, p. 6-7, fig. 3D, E. Family Basiliolidae Cooper, 1959 2010 Basiliola beecheri - Zezina, p. 1179. Genus Basiliola Dall, 1908 2015 Basiliola beecheri - Bitner, pp. 35-36, fig. 2G-H. Type species - Hemithyris beecheri Dall, 1895, by original designation of Material examined: Tonga, BORDAU 2 cruise, stn DW Dall (1908: 442) 1509, one articulated specimen (MNHN IB-2009-544); stn CP 1510, 7 articulated specimens (MNHN IB-2013-681); stn DW Basiliola lucida (Gould, 1862) 1538, 2 articulated specimens (MNHN IB-2009-545); stn CP 1539, 2 articulated specimens (MNHN IB-2009-542); stn DW 1543, one Fig. 2D-H articulated specimen (MNHN IB-2013-682); stn DW 1544, one articulated specimen (MNHN IB-2009-540); stn CP 1545, 15 arti- 1862 Rhynchonella lucida Gould, p. 120. culated specimens, one ventral and one dorsal valves (MNHN IB- 2008 Basiliola lucida - Bitner, p. 427, fig. 5A-G cum( syn.). 2009-538, IB-2013-683); stn CP 1546, 2 articulated specimens and 2009 Basiliola lucida - Bitner, p. 7, fig. 3A-C. one dorsal valve (MNHN IB-2009-541); stn DW 1548, one articu- 2010 Basiliola lucida - Zezina, p. 1179. lated specimen (MNHN IB-2013-684); stn DW 1554, one articula- 2015 Basiliola lucida - Bitner, p. 36, fig. 2C-D. ted specimen (MNHN IB-2013-685); stn DW 1555, one articulated specimen (MNHN IB-2013-686); stn CH 1557, one articulated spe- Material examined: Tonga, BORDAU 2 cruise, stn DW 1516, 4 articulated specimens